Replicated throughfall exclusion experiment in an Indonesian perhumid rainforest: wood production, litter fall and fine root growth under simulated drought.

Climate change scenarios predict increases in the frequency and duration of ENSO-related droughts for parts of South-East Asia until the end of this century exposing the remaining rainforests to increasing drought risk. A pan-tropical review of recorded drought-related tree mortalities in more than 100 monitoring plots before, during and after drought events suggested a higher drought-vulnerability of trees in South-East Asian than in Amazonian forests. Here, we present the results of a replicated (n = 3 plots) throughfall exclusion experiment in a perhumid tropical rainforest in Sulawesi, Indonesia. In this first large-scale roof experiment outside semihumid eastern Amazonia, 60% of the throughfall was displaced during the first 8 months and 80% during the subsequent 17 months, exposing the forest to severe soil desiccation for about 17 months. In the experiment's second year, wood production decreased on average by 40% with largely different responses of the tree families (ranging from -100 to +100% change). Most sensitive were trees with high radial growth rates under moist conditions. In contrast, tree height was only a secondary factor and wood specific gravity had no influence on growth sensitivity. Fine root biomass was reduced by 35% after 25 months of soil desiccation while fine root necromass increased by 250% indicating elevated fine root mortality. Cumulative aboveground litter production was not significantly reduced in this period. The trees from this Indonesian perhumid rainforest revealed similar responses of wood and litter production and root dynamics as those in two semihumid Amazonian forests subjected to experimental drought. We conclude that trees from paleo- or neotropical forests growing in semihumid or perhumid climates may not differ systematically in their growth sensitivity and vitality under sublethal drought stress. Drought vulnerability may depend more on stem cambial activity in moist periods than on tree height or wood specific gravity.

Interspecific variation in functional traits in relation to species climatic niche optima in Andean Polylepis (Rosaceae) tree species: evidence for climatic adaptations.

Plant functional traits can be genetically determined or phenotypically plastic. We assessed the degree of genetic determinism in the functional traits of Andean Polylepis tree species among 14 important traits that enable the species to withstand cold and dry conditions. We conducted a common garden experiment and related the species-specific means of the functional traits to the variables of climatic niche optima of the species (mean annual temperature and annual precipitation), deducing that if the interspecific variation in the functional trait is related to the species climatic niche optima according to the theoretically-expected pattern of climate-trait relationship, the variation of the trait must be genetically determined. In general, the traits were related either to species temperature or precipitation optima. For example, leaf size, maximum photosynthesis rate and root tip abundance were related to temperature, whereas light compensation and light saturation points were related to precipitation. Only leaf size showed a significant phylogenetic signal, indicating that most of the manifested climate-trait relationships are not caused purely by phylogeny, but are mainly a result of species specialisation along an environmental gradient. However, in many cases the relationships were rather weak. This suggests that important functional traits of Polylepis species involve both genetic and phenotypic components aiming to maximise the overall fitness of the species at high elevations.

Stem water storage in five coexisting temperate broad-leaved tree species: significance, temporal dynamics and dependence on tree functional traits.

The functional role of internal water storage is increasingly well understood in tropical trees and conifers, while temperate broad-leaved trees have only rarely been studied. We examined the magnitude and dynamics of the use of stem water reserves for transpiration in five coexisting temperate broad-leaved trees with largely different morphology and physiology (genera Fagus, Fraxinus, Tilia, Carpinus and Acer). We expected that differences in water storage patterns would mostly reflect species differences in wood anatomy (ring vs. diffuse-porous) and wood density. Sap flux density was recorded synchronously at five positions along the root-to-branch flow path of mature trees (roots, three stem positions and branches) with high temporal resolution (2 min) and related to stem radius changes recorded with electronic point dendrometers. The daily amount of stored stem water withdrawn for transpiration was estimated by comparing the integrated flow at stem base and stem top. The temporal coincidence of flows at different positions and apparent time lags were examined by cross-correlation analysis. Our results confirm that internal water stores play an important role in the four diffuse-porous species with estimated 5-12 kg day(-1) being withdrawn on average in 25-28 m tall trees representing 10-22% of daily transpiration; in contrast, only 0.5-2.0 kg day(-1) was withdrawn in ring-porous Fraxinus. Wood density had a large influence on storage; sapwood area (diffuse- vs. ring-porous) may be another influential factor but its effect was not significant. Across the five species, the length of the time lag in flow at stem top and stem base was positively related to the size of stem storage. The stem stores were mostly exhausted when the soil matrix potential dropped below -0.1 MPa and daily mean vapor pressure deficit exceeded 3-5 hPa. We conclude that stem storage is an important factor improving the water balance of diffuse-porous temperate broad-leaved trees in moist periods, while it may be of low relevance in dry periods and in ring-porous species.

Changes in wood density, wood anatomy and hydraulic properties of the xylem along the root-to-shoot flow path in tropical rainforest trees.

It is generally assumed that the largest vessels are occurring in the roots and that vessel diameters and the related hydraulic conductance in the xylem are decreasing acropetally from roots to leaves. With this study in five tree species of a perhumid tropical rainforest in Sulawesi (Indonesia), we searched for patterns in hydraulic architecture and axial conductivity along the flow path from small-diameter roots through strong roots and the trunk to distal sun-canopy twigs. Wood density differed by not more than 10% across the different flow path positions in a species, and branch and stem wood density were closely related in three of the five species. Other than wood density, the wood anatomical and xylem hydraulic traits varied in dependence on the position along the flow path, but were unrelated to wood density within a tree. In contrast to reports from conifers and certain dicotyledonous species, we found a hump-shaped variation in vessel diameter and sapwood area--specific conductivity along the flow path in all five species with a maximum in the trunk and strong roots and minima in both small roots and twigs; the vessel size depended on the diameter of the organ. This pattern might be an adaptation to the perhumid climate with a low risk of hydraulic failure. Despite a similar mean vessel diameter in small roots and twigs, the two distal organs, hydraulically weighted mean vessel diameters were on average 30% larger in small roots, resulting in ∼ 85% higher empirical and theoretical specific conductivities. Relative vessel lumen area in percent of sapwood area decreased linearly by 70% from roots to twigs, reflecting the increase in sclerenchymatic tissue and tracheids in acropetal direction in the xylem. Vessel size was more closely related to the organ diameter than to the distance along the root-to-shoot flow path. We conclude that (i) the five co-occurring tree species show convergent patterns in their hydraulic architecture despite different growth strategies, and (ii) the paradigm assuming continuous acropetal vessel tapering and decrease in specific conductance from fine roots towards distal twigs needs reconsideration.

Environmental control of daily stem growth patterns in five temperate broad-leaved tree species.

Tree ring analysis investigates growth processes at time horizons of several weeks to millennia, but lacks the detail of short-term fluctuation in cambial activity. This study used electronic high-precision dendrometry for analyzing the environmental factors controlling stem diameter variation and radial growth in daily resolution in five co-existing temperate broad-leaved tree species (genera Fraxinus, Acer, Carpinus, Tilia and Fagus) with different growth and survival strategies. Daily stem radius change (SRC(d)) was primarily influenced by the atmospheric demand for water vapor (expressed either as vapor pressure deficit (D) or relative air humidity (RH)) while rainfall, soil matrix potential, temperature and radiation were only secondary factors. SRC(d) increased linearly with increasing RH and decreasing D in all species. The positive effect of a low atmospheric water vapor demand on SRC(d) was largest in June during the period of maximal radial growth rate and persisted when observation windows of 7 or 21 days instead of 1 day were used. We found a high synchronicity in the day-to-day growth rate fluctuation among the species with increment peaks corresponding to air humidity maxima, even though the mean daily radial growth rate differed fivefold among the species. The five -species also differed in the positive slope of the growth/RH relationship with the steepest increase found in Fraxinus and the lowest in Fagus. We explain the strong positive effect of high RH and low D on radial stem increment by lowered transpiration which reduces negative pressure in the conducting system and increases turgor in the stem cambium cells, thereby favoring cell division and expansion. The results suggest that mechanistic models of tree growth need to consider the atmospheric water status in addition to the known controlling environmental factors: temperature, soil moisture and precipitation. The results further have implications for sensitivity analyses of tree growth to climatic changes.

Patterns of wood carbon dioxide efflux across a 2,000-m elevation transect in an Andean moist forest.

During a 1-year measurement period, we recorded the CO(2) efflux from stems (R(S)) and coarse woody roots (R(R)) of 13-20 common tree species at three study sites at 1,050, 1,890 and 3,050 m a.s.l. in an Andean moist forest. The objective of this work was to study elevation changes of woody tissue CO(2) efflux and the relationship to climate variation, site characteristics and growth. Furthermore, we aim to provide insights into important respiration-productivity relationships of a little studied tropical vegetation type. We expected R(S) and R(R) to vary with dry and humid season conditions. We further expected R(S) to vary more than R(R) due to a more stable soil than air temperature regime. Seasonal variation in woody tissue CO(2) efflux was indeed mainly attributable to stems. At the same time, temperature played only a small role in triggering variations in R(S). At stand level, the ratio of C release (g C m(-2) ground area year(-1)) between stems and roots varied from 4:1 at 1,050 m to 1:1 at 3,050 m, indicating the increasing prevalence of root activity at high elevations. The fraction of growth respiration from total respiration varied between 10 (3,050 m) and 14% (1,050 m) for stems and between 5 (1,050 m) and 30% (3,050 m) for roots. Our results show that respiratory activity and hence productivity is not driven by low temperatures towards higher elevations in this tropical montane forest. We suggest that future studies should examine the limitation of carbohydrate supply from leaves as a driver for the changes in respiratory activity with elevation.

Elevational change in woody tissue CO2 efflux in a tropical mountain rain forest in southern Ecuador.

Much uncertainty exists about the magnitude of woody tissue respiration and its environmental control in highly diverse tropical moist forests. In a tropical mountain rain forest in southern Ecuador, we measured the apparent diurnal gas exchange of stems and coarse roots (diameter 1-4 cm) of trees from representative families along an elevational transect with plots at 1050, 1890 and 3050 m a.s.l. Mean air temperatures were 20.8, 17.2 and 10.6 degrees C, respectively. Stem and root CO(2) efflux of 13 to 21 trees per stand from dominant families were investigated with an open gas exchange system while stand microclimate was continuously monitored. Substantial variation in respiratory activity among and within species was found at all sites. Mean daily CO(2) release rates from stems declined 6.6-fold from 1.38 micromol m(-2) s(-1) at 1050 m to 0.21 micromol m(-2) s(-1) at 3050 m. Mean daily CO(2) release from coarse roots decreased from 0.35 to 0.20 micromol m(-2) s(-1) with altitude, but the differences were not significant. There was, thus, a remarkable shift from a high ratio of stem to coarse root respiration rates at the lowest elevation to an apparent equivalence of stem and coarse root CO(2) efflux rates at the highest elevation. We conclude that stem respiration, but not root respiration, greatly decreases with elevation in this transect, coinciding with a substantial decrease in relative stem diameter increment and a large increase in fine and coarse root biomass production with elevation.

Variability in radial sap flux density patterns and sapwood area among seven co-occurring temperate broad-leaved tree species.

Forest transpiration estimates are frequently based on xylem sap flux measurements in the outer sections of the hydro-active stem sapwood. We used Granier's constant-heating technique with heating probes at various xylem depths to analyze radial patterns of sap flux density in the sapwood of seven broad-leaved tree species differing in wood density and xylem structure. Study aims were to (1) compare radial sap flux density profiles between diffuse- and ring-porous trees and (2) analyze the relationship between hydro-active sapwood area and stem diameter. In all investigated species except the diffuse-porous beech (Fagus sylvatica L.) and ring-porous ash (Fraxinus excelsior L.), sap flux density peaked at a depth of 1 to 4 cm beneath the cambium, revealing a hump-shaped curve with species-specific slopes. Beech and ash reached maximum sap flux densities immediately beneath the cambium in the youngest annual growth rings. Experiments with dyes showed that the hydro-active sapwood occupied 70 to 90% of the stem cross-sectional area in mature trees of diffuse-porous species, whereas it occupied only about 21% in ring-porous ash. Dendrochronological analyses indicated that vessels in the older sapwood may remain functional for 100 years or more in diffuse-porous species and for up to 27 years in ring-porous ash. We conclude that radial sap flux density patterns are largely dependent on tree species, which may introduce serious bias in sap-flux-derived forest transpiration estimates, if non-specific sap flux profiles are assumed.

Laser ablation-combustion-GC-IRMS--a new method for online analysis of intra-annual variation of delta13C in tree rings.

We present a new, rapid method for high-resolution online determination of delta13C in tree rings, combining laser ablation (LA), combustion (C), gas chromatography (GC) and isotope ratio mass spectrometry (IRMS) (LA-C-GC-IRMS). Sample material was extracted every 6 min with a UV-laser from a tree core, leaving 40-microm-wide holes. Ablated wood dust was combusted to CO2 at 700 degrees C, separated from other gases on a GC column and injected into an isotope ratio mass spectrometer after removal of water vapor. The measurements were calibrated against an internal and an external standard. The tree core remained intact and could be used for subsequent dendrochronological and dendrochemical analyses. Cores from two Scots pine trees (Pinus sylvestris spp. sibirica Lebed.) from central Siberia were sampled. Inter- and intra-annual patterns of delta13C in whole-wood and lignin-extracted cores were indistinguishable apart from a constant offset, suggesting that lignin extraction is unnecessary for our method. Comparison with the conventional method (microtome slicing, elemental analysis and IRMS) indicated high accuracy of the LA-C-GC-IRMS measurements. Patterns of delta13C along three parallel ablation lines on the same core showed high congruence. A conservative estimate of the precision was +/- 0.24 per thousand. Isotopic patterns of the two Scots pine trees were broadly similar, indicating a signal related to the forest stand's climate history. The maximum variation in delta13C over 22 years was about 5 per thousand, ranging from -27 to -22.3 per thousand. The most obvious pattern was a sharp decline in delta13C during latewood formation and a rapid increase with spring early growth. We conclude that the LA-C-GC-IRMS method will be useful in elucidating short-term climate effects on the delta13C signal in tree rings.